A revision of Octolepis Oliv. (Thymelaeaceae,Octolepidoideae)€¦ · species of Octolepis(with three forms) from Madagascar, named Octolepis dioicaCapuron; African Octolepisare monoecious

89

Zachary S. ROGERSMissouri Botanical Garden, P.O. Box 299, St Louis, MO 63166-0299 (USA)

[email protected]

A revision of Octolepis Oliv.(Thymelaeaceae, Octolepidoideae)

ABSTRACTA taxonomic revision of Octolepis Oliv. (Thymelaeaceae) based on morpho-logy is presented. Six species are recognized. The sole continental African spe-cies, O. casearia Oliv., is the only member of sect. Octolepis; the five otherspecies, all from Madagascar, are placed in the new sect. Dioicae. Two varie-ties are recognized for O. casearia (var. casearia and flamignii); O. decalepisGilg is placed formally into synonymy for the first time under O. casearia.Three of the five species of sect. Dioicae are described as new (O. aymoni-niana, O. ibityensis, and O. ratovosonii); O. dioica Capuron f. oblanceolataCapuron is raised to the rank of species. This revision includes full descrip-tions, illustrations of new species, identification keys, and conservation assess-ments for each taxon.

RÉSUMÉ Révision d’ Octolepis Oliv. (Thymelaeaceae, Octolepidoideae).Une révision taxonomique d’Octolepis Oliv. (Thymelaeaceae) basée sur lamorphologie est présentée. Six espèces sont reconnues. La seule espèce inféo-dée au continent africain, O. casearia Oliv., est l’unique membre de la sect.Octolepis ; les cinq autres espèces de Madagascar sont placées dans une nou-velle sect. Dioicae. Deux variétés sont reconnues au sein de O. casearia (var.casearia et flamignii) ; O. decalepis Gilg est formellement mis en synonymiesous O. casearia pour la première fois. Trois des cinq espèces de la sect.Dioicae sont nouvellement décrites (O. aymoniniana, O. ibityensis et O. rato-vosonii) ; O. dioica Capuron f. oblanceolata Capuron est élevée au rang d’es-pèce. Cette révision inclut des descriptions complètes, des illustrations desnouvelles espèces, des clés d’identification et des estimations du statut deconservation pour chaque taxon.

ADANSONIA, sér. 3 • 2005 • 27 (1) : 89-111© Publications Scientifiques du Muséum national d’Histoire naturelle, Paris.

KEY WORDSThymelaeaceae,

Octolepidoideae, Octolepis,

Africa, Madagascar,

conservation, new species.

MOTS CLÉSThymelaeaceae,

Octolepidoideae, Octolepis, Afrique,

Madagascar, conservation,

nouvelles espèces.

INTRODUCTION

Thymelaeaceae are a cosmopolitan familycomposed of about 40-45 genera and 700-800 species (MABBERLEY 1997; HERBER 2003).Molecular studies utilizing rbcL (ALVERSON et al.1998; FAY et al. 1998; SAVOLAINEN et al. 2000)and combined chloroplast and atpB sequencedata (BAYER et al. 1999) suggest that Thyme-laeaceae are one of the basal families of the orderMalvales Dumort. WURDACK & HORN (2001),on the basis of 18S rDNA, atpB and rbcLsequence data, found that Thymelaeaceae aresister to Tepuianthus Maguire & Steyerm.(= Tepuianthaceae Maguire & Steyerm.), a genusalmost entirely restricted to the “tepuis” (flat-topped sandstone mountains) of Venezuela andadjacent regions of Brazil and Colombia. HORN

(2004) provides a detailed discussion regarding therelationship of Octolepidoideae to Tepuianthus.

Based on new morphological and palynologicaldata, HERBER (2002, 2003) adjusted the limits ofthe subfamilial groups within Thymelaeaceae,resulting in the recognition of two subfamilies,Octolepidoideae Gilg (= Gonystyloideae Tiegh.+ Octolepis Oliv.; 8 genera, c. 50 spp.), andThymelaeoideae Burnett (37 genera, c. 750 spp.).Subfamily Octolepidoideae is clearly distin-guished morphologically from Thymelaeoideaeby its short or non-tubular flower, 3-12 carpellategynoecium, capsular fruit, and non-crotonoidexine pattern of the pollen grain (HERBER 2002,2003). HERBER (2003) further subdivided Octo-lepidoideae into two informally ranked groups.The Octolepis group differs from the Gonystylusgroup by straight, peltate, or reflexed (vs horse-shoe-shaped) anthers, a straight (vs contorted) style,the pubescent (vs glabrous) seed coat, striate (vsuniform) endotegmen, copious (vs absent) endo-sperm, and flat (vs thickened) cotyledons. Thegroup is composed of Octolepis, a small genus ofshrubs to medium-sized trees in continental Africaand Madagascar, and four other genera, endemicto New Caledonia and Australia (Queenslandand Northern Territory), namely Arnhemia AiryShaw (1 sp.), Deltaria Steenis (1 sp.), LethedonSpreng. (c. 15 spp.), and Solmsia Baill. (2 spp.).

Octolepis is distinguished from other genera inOctolepidoideae by a unique suite of morpholog-

ical characters, including imbricate aestivation ofthe sepals, straight filaments in flower bud, (8) 10(12) (20) stamens, basifixed anthers, and 3-6 loc-ular ovaries. Basifixed anthers are probably asynapomorphy of the genus according to HERBER

(2003). The taxonomy of Octolepis has never been

revised and species limits within the genus havebeen particularly unclear. This revision specifi-cally addresses the taxonomic problems associ-ated with the continental African taxa (theOctolepis casearia species complex) and theMalagasy taxa (the Octolepis dioica complex).

TAXONOMIC HISTORY OF OCTOLEPIS

OLIVER (1865) first described Octolepis basedon a single species from Nigeria (O. caseariaOliv.). Over the course of the next 50 years, sev-eral authors published seven additional names inOctolepis for taxa with types from the adjacenttropical African countries of Cameroon,Democratic Republic of the Congo (Kinshasa),Gabon, Ivory Coast, and Liberia. GILG (1899)was responsible for publishing four of thosenames, each based on only one or two specimens.He distinguished his species by single characters,such as leaf shape and size, acumen length, num-ber of secondary veins, and flower meristicity (4-vs 5-merous flowers). The large amount of mor-phological plasticity in these characters, in addi-tion to the lack of sufficient material at the timethe species were described, led later authors(AYMONIN 1966a, b; ROBYNS 1975) to considernearly all species of the O. casearia complex to besynonymous with O. casearia. Most authors haverecognized a second species of Octolepis in Africa,(O. decalepis Gilg) because of its (almost) always5-merous flowers.

CAPURON (1963) described a single dioeciousspecies of Octolepis (with three forms) fromMadagascar, named Octolepis dioica Capuron;African Octolepis are monoecious and have bisex-ual flowers. The genus has rarely been mentionedin the literature since the description of O. dioica.

Dioecy, or at least partial dioecy (e.g., gyno-dioecy or functional dioecy), has been recordedin both subfamilies of Thymelaeaceae, and in

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90 ADANSONIA, sér. 3 • 2005 • 27 (1)

14 genera, including Lethedon and Solmsia (HERBER

2003). These two genera may also be the mostclosely-related genera to Octolepis based on mor-phology (HERBER 2002, 2003) and a preliminarymolecular analysis (WURDACK unpubl. data).

MATERIALS AND METHODS

This revision is based on morphological datagathered from herbarium specimens examinedfrom the following institutions: A, B, BM, BR, F,G, GH, K, MA, MO, NY, P, TAN, TEF, US,WAG. Because Malagasy material was poorlyknown and under represented in herbaria, severalmonths were spent observing the species in situand collecting additional material. After two peri-ods of collection at different times of the year(January-March 2003 and March-July 2004), fer-tile material was collected for representatives offour of the five Malagasy species recognized inthe revision. I was unable to find O. aymoninianaZ.S.Rogers in the field.

Descriptive terminology follows STEARN

(1992). Measurements are taken from fertilespecimens. Flower width was determined by mea-suring the distance between opposing sepal tipslaid flat. The number of petals in the flowers ofOctolepis has been interpreted differently byauthors. This confusion is caused by whether ornot the petals are truly divided almost entirely tothe base, making them bifid and typically doublethe number of sepals, or not divided at all, andequal to the number of sepals. I have found vari-ous degrees of fusion at the base of the petal lobesin both African and Malagasy species, but havemost often observed fusion of two adjacent lobesc. 1 mm from the receptacle. Petals are describedas bifid in the remainder of the paper.

Keys are intended for use with fertile material.Several Malagasy collections made from low andmid-elevation forests, areas where Octolepis hasrarely been recorded, exhibit somewhat differentmorphology from the species enumerated below.This material has been excluded from thedescriptions and specimen citation, but thesespecimens are discussed under either O. dioicaand O. oblanceolata, where they would be identi-fied using the key to species.

Complete collection data for cited exsiccataeand photos of types and other representativespecimens are posted on the W3 TROPICOSdatabase at: http://mobot.mobot.org/W3T/Search/vast.html/. Coordinates and elevations ofcollecting localities in mainland Africa wereassigned post-facto, when necessary, using severalon-line gazetteers and paper maps; Malagasy col-lections lacking coordinates and elevations wereassigned with the “Gazetteer to MalagasyBotanical Collecting Localities” (SCHATZ &LESCOT 2005: http://www.mobot.org/MOBOT/Research/madagascar/gazetteer/). Post-factocoordinates and elevations are enclosed by squarebrackets in the text. The distribution maps werecreated using ArcView GIS (version 3.2a); dis-tributions of the Malagasy species are mappedover the five simplified bioclimatic zones ofMadagascar (SCHATZ 2000 following CORNET

1974). Conservation status for each species isprovisionally assigned based on the Red ListCategories and Criteria version 3.1 (IUCN 2001).

The species concept and criteria (i.e. the waythe species are recognized) follow those proposedin ROGERS (2004). Unlike that study, however,infraspecific taxa (viz. two varieties) are recog-nized by a single consistent morphological char-acter difference coupled with a slightly allopatricdistribution.

SYSTEMATICS

OCTOLEPIS Oliv.

J. Linn. Soc. Bot. 8: 161, t. 12 (1865).

Makokoa Baill., Bull. Mens. Soc. Linn. Paris 1: 619(1886). — Type: Makokoa congolana Baill.

TYPE. — Octolepis casearia Oliv.

Shrubs or trees; plants monoecious (Africa) ordioecious (Madagascar); bark longitudinallystriate, sometimes lenticellate. Leaves alternate oropposite, often discolorous, caducous or persis-tent on mature stems, estipulate; leaf bladesglabrous adaxially, sparsely covered with incon-spicuous adpressed indument abaxially orglabrescent (trichomes c. 0.5-1 mm long), margin

Revision of Octolepis (Thymelaeaceae)

91ADANSONIA, sér. 3 • 2005 • 27 (1)

entire, ± revolute; midrib grooved adaxially,raised abaxially; venation brochidodromous; finevenation reticulate; leaf buds conduplicatelyfolded, covered with dense indument; petiolesgrooved adaxially, articulate, drying darker thanmidrib. Inflorescences axillary and fasciculate(Malagasy species), or lateral (or sublateral) andborne on short woody compact racemose axes(African species); inflorescence bracts absent;pedicels articulate. Flowers white, small, bisexualor unisexual; staminate and pistillate flowers ofsimilar size and shape; floral tube absent; sepals 4-6,imbricate, reflexed or spreading, rarely erect, per-sistent in fruit; petals scale-like, 4 or 5 (10), bifid,divided nearly to base, adnate to receptacle, per-sistent in fruit; stamens twice the number ofpetals, (8) 10 (12) (20), straight in bud, ± insertedin a single whorl, persistent in fruit; anthersoblong, introrse, basifixed. Staminate flowerswith a minute pistillode; pistillode obscured bymany erect receptacle trichomes. Pistillate flowerswith (8) 10 (12) (20) persistent staminodes;rudimentary anthers globose or undifferentiated;subgynoecial disc absent; gynoecium 3-6-carpel-late, sessile; carpels uniovulate; ovary superior,usually one to several locules aborting, denselypubescent, surrounded by many erect receptacletrichomes; ovules minute, anatropous, pendu-lous; style terminal, straight, persistent; stigmaglobose, lobed, papilliate. Fruits green-yellow orwhite (O. casearia), loculicidal capsules, sub-spheroidal to ovoid-pyramidal, developed carpels

producing longitudinal lines of dehiscence; dehis-cence lines depressed or ± plane, often bicarinateon both sides of line; pericarp dry and thin, orfleshy and thicker. Seeds brown or black, ovoid,triangular or circular in cross section, edgesrounded, adaxial side often costate, curved, abax-ial side costate, other sides curved or ± flat; seedcoat crustaceous, thin, outer epidermis puberu-lent, indument red-orange, denser near poles,inner epidermis finely striate; endosperm copi-ous; cotyledons longitudinally curved, thin(c. 0.5-1.5 mm thick).

INFRAGENERIC CLASSIFICATION OF OCTOLEPIS

Octolepis casearia, the continental Africanspecies, differs significantly from the Malagasytaxa by a number of vegetative, floral, and fruitcharacters (Table 1). The petals of O. casearia(Fig. 1A, B) have an obvious papilliate surfacesculpturing. In O. dioica (Fig. 1C), the petals arealso papilliate, but the sculpture pattern is rarelyevident because of the dense indument coveringthe lobes. Petal indument in O. casearia is alwaystomentose or tomentose-strigose, usuallyconcentrated along the margin and apex of thelobes, and the trichomes are not oriented in aparticular direction. The petal indument ofO. dioica is representative of the other Malagasyspecies and is characterized as strigose (nevertomentose) on both surfaces with the individualtrichomes always oriented toward the receptacle(receptacle located just outside the bottom of the

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92 ADANSONIA, sér. 3 • 2005 • 27 (1)

TABLE 1. — Morphological differences in Octolepis between sect. Octolepis and sect. Dioicae.

Character Sect. Octolepis Sect. Dioicae

Sexual system monoecious dioeciousHabit shrubs 2-4 m tall, rarely trees to 7 m tall trees to 20 m tallLeaves persistent caducousInflorescence position lateral or sublateral, borne on foliated stems axillary, usually borne on defoliated

stemsInflorescence type compact racemose axes (often fasciculate

elongating into spurs)Petal orientation erect adpressed to sepalsPetal lobe shape broadly oblong narrowly triangularLobe length/width ratio c. 1-1.5:1 c. 4-7:1Petal lobe apex broadly truncate or rounded acute, rounded at tipPetal indument tomentose or tomentose-strigose strigosePetal trichome orientation no particular direction toward receptaclePericarp dry, thin, smooth fleshy, thick, rugose or rugulose

frame in Figure 1C). Populations of African andMalagasy Octolepis are separated geographicallyby c. 3000 km.

In addition to the substantial morphologicaland geographical differences, these two groupsform distinct monophyletic lineages. CombinedrbcL and trn-F sequence data taken from the fiverecognized Malagasy species, both varieties ofO. casearia, and a sample of O. decalepis (sensuGilg), indicate that each group is monophyleticwith 99% bootstrap support (WURDACK unpubl.data).

Based on these strong lines of supporting evi-dence, the following infrageneric classificationfor Octolepis is proposed, to stress the significantdifferences between these two groups. Two sec-tions are recognized here: sect. Octolepis (thesole continental African species, O. casearia) andsect. Dioicae Z.S.Rogers (the five Malagasyspecies).

Octolepis Oliv. sect. Octolepis

TYPE. — Octolepis casearia Oliv.

Monoecious shrubs, rarely small trees; barkusually lenticellate. Leaves alternate, persistent onmature stems, apex usually not aborting.Inflorescences lateral or sublateral, borne on thefoliated portion of the stems; axes compact, race-mose, elongating with up to c. 30 flowers perinflorescence. Flowers bisexual; sepals spreadingor reflexed; petals 4 or 5, rarely 6; petal lobesbroadly oblong, length/width ratio c. 1-1.5:1,erect, apex truncate or rounded; stamens 8 or 10.Fruits white, dry, dehiscence lines ± plane, notcarinate on side of dehiscence lines; pericarp < 2 mmthick, smooth when dry. Seeds with cotyledonsc. 0.5(-1) mm thick [plants of continental Africa].

Octolepis Oliv. sect. Dioicae Z.S.Rogers, sect.nov.

TYPE. — Octolepis dioica Capuron.

Haec sectio a sectione Octolepide speciebus monoiciis(nec diociis), habitu plerumque fruticoso (nec arborescente),


93ADANSONIA, sér. 3 • 2005 • 27 (1)

A

B

C

L

S

1000 μm

1000 μm

100 μm

FIG. 1. — SEM microphotograph of the petals in Octolepis:A, O. casearia (sect. Octolepis), adaxial surface of sepals (S)and abaxial surface of petal lobes (L). The arrow indicates thepoint where two erect lobes meet to form a single deeply divid-ed bifid petal. The stamens located just above petal lobes areout of frame; B, papilliate abaxial surface of the petal shown in Aat higher magnification; C, O. dioica (sect. Dioicae), adaxial sur-face of a bifid petal. A, B, Breteler et al. 9450, WAG pickledsample; C, Razakamalala 748, MO pickled sample.

foliis persistentibus (nec caducis), inflorescentiis lateral-ibus (nec terminalibus), petalis lobis late oblongis (nectriangularibus) et pericarpio tenui pagina laevi (neccrasso pagina plus minusve rugosa) differt.

Dioecious trees; bark rarely lenticellate. Leavesalternate to opposite, caducous on mature stems,apex often aborting. Inflorescences axillary,fasciculate; fascicles 1 to several-flowered,usually borne on defoliated portion of stems.Flowers unisexual; sepals spreading or erect;petals 4-6 (10); petal lobes narrowly triangular,

length/width ratio c. 4-7:1, spreading, adpressedto sepals, apex acute, rounded at the tip.Staminate flowers with (8) 10 (12) (20) stamens;pistillode minute, obscured by many receptacletrichomes. Pistillate flowers with (8) 10 (12) (20)staminodes. Fruits green-yellow, fleshy, oftenbicostate on both sides of dehiscence lines;per icarp 2-8 mm thick, rugose or rugulose,often pitted, rarely smooth when dry. Seedswith cotyledons 1-1.5 mm thick [plants ofMadagascar].

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94 ADANSONIA, sér. 3 • 2005 • 27 (1)

Key to the species of Octolepis(Leaves of Malagasy species illustrated in Figure 2)

1. Monoecious shrubs, rarely small trees; petal lobes broadly oblong (length/width ratio 1-1.5:1), apex truncateor rounded; fruits dry; [plants of Africa] ................................................................................ 2. O. casearia

1’. Dioecious trees; petal lobes narrowly triangular (length/width ratio 4-7:1), apex acute, rounded at the tip;fruits fleshy; [plants of Madagascar] .......................................................................................................... 2

2. Leaf blades chartaceous, apex attenuate-acuminate; secondary veins 15-25 pairs, closely-spaced; pedicels 0.3-0.5 mm in diam.; receptacle trichomes surrounding the gynoecium 2.5-3.0 mm long in pistillate flowers .......................................................................................................................................... 1. O. aymoniniana

2’. Leaf blades coriaceous or coriaceous-chartaceous, apex usually slightly emarginate to retuse, sometimesacute, rounded, or obtuse, rarely short cuspidate; secondary veins rarely up to 12 pairs, sparsely-spaced;pedicels more than 0.5 mm in diam.; receptacle trichomes surrounding the gynoecium up to 2.5 mm longin pistillate flowers .................................................................................................................................... 3

3. Petioles 8-16 mm long; pedicels (in flower or fruit) 1.6-2.5 cm long, (1.5-)2-3 mm in diam.; sepals 5-9 mmwide; bidif petals 10 (i.e. 20 lobes); stamens and staminodes 20 ...................................... 6. O. ratovosonii

3’. Petioles up to 8 mm long; pedicels (in flower or fruit) ≤ 1.5 cm long, up to 1 mm in diam.; sepals 2-4 mmwide; bifid petals 4-6 (i.e. 8-12 lobes); stamens and staminodes (8) 10 (12) .............................................. 4

4. Widest leaf blades 1-1.4 cm wide; pedicels less than 2 mm long; flowers 4-6 mm wide; largest fruits 1.6 cmlong, dehiscence lines 1 or 2, rarely 3 .................................................................................. 4. O. ibityensis

4’. Widest leaf blades (1.6-)2-4 cm wide; pedicels greater than 4 mm long; flowers more than 8 mm wide;largest fruits 2-3 cm long, dehiscence lines 4 or 5 .................................................................................... 5

5. Longest leaf blades 2.8-6.0(-6.5) cm long, c. 1-2.5 times longer than wide; leaf margin flat; flowers 8-11 mmwide ........................................................................................................................................ 3. O. dioica

5’. Longest leaf blades (7.0-)7.4-12.4 cm long, c. 3-6 times longer than wide; leaf margin undulate; flowers 11-13 mm wide .................................................................................................................. 5. O. oblanceolata

1. Octolepis aymoniniana Z.S.Rogers, sp. nov.

Haec species a speciebus aliis Octolepidis laminafoliari tenui marginibus haud revolutis, venis secon-dariis numerosis (15 ad 25), pedicellis filamento-s i s 0 .3 -0 .5 mm in d iamet ro e t t r i chomat ibu snumerosissimis 2.5-3 mm longis gynoecium cingen-tibus differt.

TYPUS. — Kotozafy & Randriamanantena 355,Madagascar, Prov. Fianarantsoa, Ranomafana ParcNational (parcelle 1), à Antara, à 7 km au N de la villed’Ambatolahy, c. 900 m, [21°14’S, 47°26’E], 19-21 Oct. 1993, fl. (holo-, MO!; iso-, TAN!, WAG!).

Treelets to 2 m tall; young stems denselystrigose-tomentose. Leaves alternate to subop-posite; blades broadly elliptic or elliptic-oblong,c. 6.1-9.7 × 1.7-2.5 cm, length/width ratio c. 3-4:1, chartaceous, surfaces ± discolorous, adaxialsurface glabrous, abaxial surface lighter green,sparsely strigose or glabrescent, apex attenuate-acuminate (rarely subacute), acumen up to9 mm long, margin sparsely strigose or glabres-cent, slightly undulate, base short attenuate orcuneate; midrib strigose; venation raised onboth surfaces, densely congested, secondary

veins 15-25 pairs, closely-spaced, angle of diver-gence from midrib 50-65°, submarginal veinloop 1-2 mm from margin; intersecondariesnearly parallel, closely-spaced; petioles 5-7 mmlong, c. 1 mm in diam., densely strigose-tomen-tose or glabrescent. Inflorescences axillary, borneon defoliated portion of stem; fascicles 1-4-flowered, 1 or 2 flowers persistent; pedicelsc. 8-14 mm long, filamentous (0.3-0.5 mm indiam.), glabrous, base strigose. Flower buds3-3.5 × 3 mm, sparsely strigose or glabrescent,surface smooth. Flowers 9-11 mm wide; sepals5, ovate or oblong, 3.6-4.5 × 1.7-2.2 mm, coria-ceous, sparsely strigose or glabrescent on bothsurfaces, apex acute, strigose-tomentose, mar-gins tomentose or glabrescent; petals 5, denselystrigose on both surfaces; petal lobes narrowlytriangular, each lobe 3.7-4.3 × 0.5-0.7 mm,length/width ratio c. 6-7:1, coriaceous, apex

acute. Staminate flowers not seen. Pistillateflowers with 10 staminodes, each 1-1.2 mmlong, 0.1-0.2 mm wide (at base), rugulose,glabrous; rudimentary anthers globose; ovary0.8-1.4 × 1.1-1.8 mm, surrounded by hundredsof erect receptacle trichomes, each 2.5-3 mmlong; style 3.1-3.8 × 0.2-0.4 mm, glabrescent,base strigose. Fruits not seen. — Fig. 3.

DISTRIBUTION AND PHENOLOGY. — Octolepisaymoniniana, known only from the type collec-tion, was made in dense, humid, primary forestwithin Ranomafana National Park at 900 m ele-vation (Fig. 7). The species was collected inflower in October.

VERNACULAR NAMES. — None available.CONSERVATION STATUS. — Octolepis aymonini-

ana is known from a single collection made in1993. Despite a thorough search of the type


95ADANSONIA, sér. 3 • 2005 • 27 (1)

A

B

C

DF

G

H

E

1 cm

FIG. 2. — Leaf variation in the Malagasy species of Octolepis (abaxial leaf surface): A, B, O. dioica Capuron; C, D, O. oblanceolata(Capuron) Z.S.Rogers; E, O. ibityensis Z.S.Rogers; F, O. aymoniniana Z.S.Rogers; G, H, O. ratovosonii Z.S.Rogers. A, Rogers &Razakamalala 46; B, Service Forestier 6028; C, Réserves Naturelles 10459, type collection; D, Randriamampionona 1308;E, Randrianaivo et al. 8, type collection; F, Kotozafy & Randriamanantena 355, type collection; G, Rogers et al. 165, type collection;H, Ratovoson et al. 247, an exceptionally large leaf.

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96 ADANSONIA, sér. 3 • 2005 • 27 (1)

A

B

C

D

1 cm

1 cm

1 cm

FIG. 3. — Octolepis aymoniniana Z.S. Rogers, sp. nov.: A, habit of pistillate individual; B, base of inflorescence and leaf scar;C, pistillate flower (note ovary surrounded by hundreds of long, erect, filamentous receptacle trichomes); D, relationship betweensepals, bifid petals, and staminodes. Kotozafy & Randriamanantena 355 (TAN, type).

locality accompanied by the original collector(KOTOZAFY), I was unable to locate other individ-uals in 2003. The site falls within the currentboundaries of Ranomafana National Park.Vegetation at the locality remains in good condi-tion, so it is likely that O. aymoniniana stilloccurs there. The species is assigned a provisionalstatus of Vulnerable (VU B1ab+B2ab).

Octolepis aymoniniana is distinguished by thehighly congested venation consisting of 15-25 pairs of closely-spaced secondary veins withmany intersecondaries, and the small, narrow,elongated areolae. The species differs markedlyfrom other members of the genus by its filamen-tous pedicels (c. 0.3-0.5 mm in diam.) and by thehundreds of long receptacle trichomes (2.5-3 mmlong) surrounding the gynoecium in pistillateflowers. Vegetatively, the species can be recog-nized from all other Malagasy Octolepis by thethin, chartaceous leaf blades with attenuate-acuminate (or infrequently subacute) apices andnon-revolute margins.

Octolepis aymoniniana is named in honor ofGérard-Guy AYMONIN, whose lifetime study ofpaleotropical Thymelaeaceae has been a greatcontribution to our knowledge of the family.

2. Octolepis casearia Oliv.

J. Linn. Soc. Bot. 8: 161, t. 12. (1865). — Type:Mann 2306, Nigeria, State Cross River, River OldCalabar, [4°54’N, 8°16’E], Feb. 1863, fl. (lecto-, K!,designated by AYMONIN 1966a).

Shrubs 2-3 m tall, rarely small trees to 7 m tall;bark usually lenticellate; stems with internodes0.9-6.1 cm long; young stems densely to sparselystrigose-tomentose. Leaves alternate, bladesbroadly obovate or elliptic, c. 5-31 × 3.5-10.5 cm, length/width ratio c. 2.5-4:1, coriaceousor rarely subcoriaceous, sometimes bullate, sur-faces usually discolorous, adaxial surfaceglabrous, abaxial surface usually lighter green,glabrescent or sparsely strigose, trichomes c. 0.3-0.5 mm long, apex acuminate or acute, acumenup to 2.2 cm long, margin revolute, glabrescentor sparsely strigose, base cuneate or short attenu-ate; midrib strongly raised on abaxial surface,

sparsely to moderately strigose; venation stronglyraised on both surfaces, secondary veins c. 6-13 pairs, usually strongly arched toward apex,angle of divergence from midrib 55-75°, sub-marginal loop c. 4-11 mm from margin; petiolesc. 4-10 mm long, strigose-tomentose. Inflo-rescences lateral or sublateral, borne on foliatedportion of the stem; axes to 6.7 mm long, c. 2 mmwide, with up to c. 30 flowers per inflorescence,densely strigose-tomentose; pedicels c. 3-8 mmlong, strigose-tomentose. Flowers 6-11 mm wide;sepals 4 or 5, rarely 6, ovate or subtriangular, 2.5-6 × 1.3-3.1 mm, reflexed or spreading, coriaceousto subcoriaceous, strigose-tomentose on both sur-faces, margins tomentose, apex acute; petals 4 or 5,rarely 6, erect, glabrescent or tomentose-strigoseon both surfaces; each petal lobe broadly oblong,1.1-3 × 0.5-1.4 mm, length/width ratio c. 1-1.5:1, coriaceous to subcoriaceous, apex truncateor rounded, densely tomentose, margin usuallydensely tomentose; stamens 8 or 10, rarely 12; fil-aments 1.4-5.5 × 0.2-0.3 mm, strigose-tomen-tose or glabrescent; anthers 0.7-1 × 0.3-0.5 mm;ovary subspheroidal, 0.9-1.5 × 0.8-1.3 mm,densely strigose-hirsute, surrounded by manyerect receptacle trichomes, each c. 0.5 mm long;style 1.1-1.6 × 0.3-0.5 mm, strigose. Fruitswhite, ovoid-suborbicular, 1-1.5 × 1-1.3 cm;dehiscence lines 2-5 (i.e. 2-5 carpels develop),plane or slightly raised; pericarp dry, 1-2 mmthick, smooth, glabrescent or sparsely strigose, orcovered with dense persistent indument. Seedsdark brown or black, 8-13 × 5-8 mm, puberu-lent, indument denser near poles and on adaxialsurface, trichomes c. 0.2-0.5 mm long, erect oradpressed.

TYPIFICATION . — In the protologue ofOctolepis casearia (OLIVER 1865), three collec-tions were cited (Mann s.n., Sept. 1862; Manns.n., Feb. 1863; and Thomson s.n.). No specimenscollected by THOMSON have been located, buttwo numbered specimens collected by MANN aredeposited in the Kew herbarium, which corre-spond to the dates cited in the protologue (Manns.n., Sept. 1862 [= Mann 1815] and Mann s.n.,Feb. 1863 [= Mann 2306]). These collectionsundoubtedly represent the two specimens men-tioned by OLIVER (1865). In the Thymelaeaceae


97ADANSONIA, sér. 3 • 2005 • 27 (1)

treatments appearing in the Flore du Gabon(AYMONIN 1966a: 46) and Flore du Cameroun(AYMONIN 1966b: 80), only one of these syntypes(Mann 2306) was mentioned as the type ofO. casearia, thereby effectively lectotypifying thename according to Article 7.11 of the Code(GREUTER et al. 2000).

Octolepis casearia is easily recognized from allother members of the genus by its bisexual flow-ers, broadly oblong petal lobes with truncate orrounded apices, dry fruit with a thin pericarp,and its central and west African distribution.

Octolepis casearia is circumscribed here toinclude all previously described continentalAfrican taxa. Earlier authors have also recognizeda broadly circumscribed O. casearia, due to theoverlapping, continuous morphological variationwithin characters used to distinguish the species.Here, O. decalepis Gilg, previously segregated bysome because of its (almost) always 5-merousflowers, is also included within O. casearia.

Flower meristicity is rather variable geographi-cally within Octolepis casearia and inconsistentamong specimens, sometimes even for collections

made from the same locality or even those takenfrom the single plant (meristicity is much morevariable within the Malagasy species). Octolepiscasearia has 5-merous flowers in Ivory Coast andalmost always in Liberia, 4-merous flowers inNigeria, 4- or 5-merous flowers in Cameroon andGabon, and 5-merous flowers in the Republic ofthe Congo (Brazzaville).

In contrast, variation in fruit morphologyclearly delimits two distinct groups of specimensof Octolepis casearia. One group with denselypubescent fruits, includes the type of O. flamigniiDe Wild., and occurs in the DemocraticRepublic of the Congo (Kinshasa), southeasternGabon, and the Republic of the Congo (Brazza-ville), while a second group with glabrescent orsparsely strigose fruits, corresponds to O. caseariavar. casearia, and grows in Cameroon, IvoryCoast, Gabon, Liberia, Nigeria, and the Republicof the Congo (Brazzaville). These two groups areslightly allopatric, but some populations comewithin c. 60 km of each other in Gabon and theDemocratic Republic of the Congo (Fig. 4).These two distinct groups are recognized asvarieties of O. casearia.

Rogers Z.S.

98 ADANSONIA, sér. 3 • 2005 • 27 (1)

Key to the varieties of Octolepis casearia

— Fruits glabrescent or sparsely strigose; [plants of Cameroon, Ivory Coast, Gabon, Liberia, Nigeria, andRepublic of the Congo (Brazzaville)] .................................................................................. 2a. var. casearia

— Fruits densely pubescent, surface obscured by indument; [plants of Democratic Republic of the Congo(Kinshasa), southeastern Gabon, and Republic of the Congo (Brazzaville)] .................... 2b. var. flamignii

2a. Octolepis casearia Oliv. var. casearia

Makokoa congolana Baill., Bull. Mens. Soc. Linn. Paris1: 619 (1886). — Type: Thollon 343, Gabon,Ogooué, “mission P. Savorgnan de Brazza”, 6 Dec.1883, fl. (holo-, P!; iso-, P[3]!). — Octolepis congo-lana (Baill.) Warb., Engl. & Prantl, Nat.Pflanzenfam. 3, 6a: 56 (1893).

Octolepis decalepis Gilg, Bot. Jahrb. Syst. 28: 142(1899). — Type: Dinklage 1741, Liberia, CountyGrand Bassa, Fishtown, [5°52’N, 10°03’W], fl.(holo-, B!; iso-, A!, B[2]!, MO!); syn. nov.

Octolepis dinklagei Gilg, Bot. Jahrb. Syst. 28: 143(1899). — Type: Dinklage 177, Cameroon, Prov.Sud, Ebea-Falle près Kribi, [2°57’N, 9°55’E],Oct. 1889, fl., fr. (lecto-, K!, here designated;iso-, S!).

Octolepis macrophylla Gilg, Bot. Jahrb. Syst. 28: 144(1899). — Type: Staudt 608, Cameroon, Prov.Sud-Ouest, Station Johann-Albrechtshohe [= Kumba],Urwaldgebiet, [4°38’N, 9°25’E], 11 Feb. 1896, fl.(lecto-, G!, here designated; iso-, A!).

Octolepis nodosericea Gilg, Bot. Jahrb. Syst. 28: 143(1899). — Type: Bos 6337, Cameroon, Prov. Sud,c. 60 km N of Kribi, riverine forest of Bivouba River,3°19’N, 10°06’E, 13 Feb. 1970, fl., fr. (neo-, WAG-0131161!, here designated; iso-, K!, WAG[2]!).

Octolepis pierreana Gilg ex Engl., in obs., Engl., Veg.Erde 9, Pflanzenw. Afr. 628 (1921). — Type:Breteler, Jongkind & Wieringa 11379, Gabon, Prov.Ogooué-Ivindo, between Ndjolé and bridge cross-ing Ogooué River, [00°06’S, 11°25’E], 30 July1992, fl., fr. (neo-, WAG-0131152!, here desig-nated; iso-, G!, WAG!).

Shrubs to 3 m tall, rarely trees to 6 m tall;young stems densely to sparsely strigose-tomen-tose. Leaf blades c. 6.6-31 × 3-10.5 cm, some-times bullate, apex acuminate or acute, acumenup to 2.2 cm long. Inflorescence axes to 6.7 mmlong, with up to c. 30 flowers per inflorescence;pedicels 4-7 mm long. Flowers with 4 or 5 (6)sepals. Fruits glabrescent or sparsely strigose; tri-chomes up to 0.8 mm long.

Refer to OLIVER (1865: pl. 12) for an illustra-tion of O. casearia var. casearia.

DISTRIBUTION AND PHENOLOGY. — Octolepiscasearia var. casearia is widely distributed fromLiberia to the Democratic Republic of the Congo(Kinshasa) from 0-760 m elevation (Fig. 4); onecollection (Thomas & Etuge 5906) from Came-roon reported an elevation of 1000-1300 m.

Note the large geographic disjunction betweenpopulations growing in the western portionof the Ivory Coast and those in south easternNigeria. The variety is frequently found in pri-mary and secondary forests. Individuals typicallygrow on sandy substrates and along river banks.Octolepis casearia var. casearia flowers and fruitsyear round. Flowers and fruits are often found onthe same plant.

VERNACULAR NAMES. — Luvaka (Hawner 39).CONSERVATION STATUS. — Based on available

label data, Octolepis casearia var. casearia appearsto be relatively common in primary and sec-ondary forests in central and western Africa. Thevariety has been collected from several protectedareas in a number of countries: Cameroon(Ejagham Forest Reserve; Korup National Park;Southern Bakundu Forest Reserve), Gabon


99ADANSONIA, sér. 3 • 2005 • 27 (1)

10°W 0 10°E 20°E

10°N

0

10°S

1000 km

FIG. 4. — Distribution map of Octolepis casearia Oliv. var. casearia (▲) and O. c. var. flamignii (De Wild.) Z.S.Rogers (●). Outlinescorrespond to country boundaries in central and western Africa.

(Lopé-Okanda Reserve; Plateaux Betéké NationalPark), Ivory Coast (Taï National Park), Liberia(Gola National Forest). Octolepis casearia var.casearia should be considered a taxon of LeastConcern (LC).

TYPIFICATION . — The type material ofOctolepis dinklagei (Dinklage 177) deposited at Bis believed to have been destroyed (R. VOGT pers.comm.). The duplicate of the type collectiondeposited at K is designated as the lectotype.

GILG (1899) cited two syntypes for Octolepismacrophylla (Staudt 608; Staudt 838). No dupli-cates of either collection are extant at B. Twoduplicates of Staudt 608, one each at A and G,have been located; the G sheet is the morecomplete specimen and is selected as the lecto-type.

Zenker 823, a collection made near Kribi,Cameroon, was originally cited as the type ofOctolepis nodosericea. No duplicates of the collec-tion have been located and the holotype thatwould have been deposited at B may have beendestroyed (R. VOGT pers. comm.). In accordancewith Article 9.6 of the Code (GREUTER et al.2000), a neotype must therefore be chosen. AsO. nodosericea was not illustrated in the proto-logue, another specimen must be selected. Bos6337 does not conflict with the description andwas made near the type locality. The WAG dupli-cate with the accession number 0131161 isselected as the neotype.

The validly published name Octolepis pierreanaGilg ex Engl. (ENGLER 1921) was distinguishedfrom O. macrophylla Gilg by its more lanceolateleaf blade with a longer acuminate apex; thespecies was noted to be from Gabon in the proto-logue. No cited specimens or illustrations accom-panied the description and original material GILG

may have used to describe the species has notbeen found, making it necessary to designate aneotype. Breteler et al. 11379 (WAG-0131152)has more lanceolate leaves than those found onthe lectotype of O. macrophylla and is chosen asthe neotype.

Octolepis casearia var. casearia is easily distin-guished from var. flamignii by the glabrescent orsparsely strigose (vs densely pubescent) fruits.

SELECTED MATERIAL EXAMINED (1 collection cited foreach major political division below country; all of the124 examined collections are available on the W3 TROPI-COS database; every georeferenced collection is plottedin Figure 4). — CAMEROON: Prov. Centre: Breteler2659, Oveng village, 27 km from Sangmélima, alongroad to Yaoundé, [3°42’N, 11°22’E], 20 Mar. 1962, fl.(A!, WAG[2]!). — Prov. Littoral: Mildbraed 8227,Douala [= Kribi], [3°45’N, 9°57’E], Feb. 1914, fl. (K!).— Prov. Sud: Bos 7022, 30 km S of Kribi, Mbodé bay,[2°45’N, 9°53’E], 7 July 1970, fl., fr. (BR!, K!, MO!).— Prov. Sud-Ouest: Thomas et al. 7531, KorupNational Park, sandy area between Baro and Ikenge vil-lages, 250 m, [5°15’N, 9°09’E], 1 Apr. 1988, fl., fr.(MO!, WAG!).GABON: Prov. Estuaire: De Wilde et al. 8826, Barragede Kinguéle, 100 m, 0°28’N, 10°17’E, 19 Nov. 1986,fl., fr. (K!, MA!, MO!, WAG[4]!). — Prov. Moyen-Ogooué: Wieringa et al. 4066, 34 km on road Alèmbéto Lopé, 0°03’35”S, 11°10’20”E, 11 Jan. 2001, fr.(WAG[2]!). — Prov. Ngounie: Wieringa et al. 2991,Fougamou, 10 km on forestry road following BendoloRiver, 180 m, 1°12’35”S, 10°31’06”E, 30 Oct. 1994,fl., fr. (WAG[2]!). — Prov. Nyanga: Le Testu 1844,Tchibanga, région du Nyanga, [2°51’S, 11°02’E],10 Nov. 1914, fl. (A!, BM!). — Prov. Ogooué-Ivindo:Breteler et al. 11379, between Ndjolé and bridge cross-ing Ogooué River, 0°06’S, 11°25’E, 8 May 1992, fl.,fr. (G!, WAG[2]!). — Prov. Ogooué-Maritime: Breteleret al. 9450, Rabi, 1°55’S, 9°50’E, 24 Mar. 1990, fl., fr.(WAG[2]!).IVORY COAST: Region Bas-Sassandra: Breteler 7367,c. 13 km NW of Tabou, [4°30’N, 7°24’W], 12 Apr.1974, fl. (BR!, MO!, WAG[2]!). — Region Dix-HuitMontagnes: Aké Assi 17804, Parc National de Taï,[5°45’N, 7°05’W], 15 Dec. 1987, fl. (G[2]!, MO!).LIBERIA: County Bomi: Stoop 201A, Bomi Hills,Gola National Forest, [6°56’N, 10°45’W], 24 Aug.1970, fr. (WAG!). — County Grand Bassa: Dinklage2044, Grand Bassa, [5°52’N, 10°03’W], fl., fr. (A!,K[2]!, MO!). — County Grand Gedeh: Jansen 1281,10 miles from Tchien, along road to Cape Palmas,[6°04’N, 8°08’W], 22 Jan. 1969, f l . (MO!,WAG[2]!). — County Montserrado: Dinklage 2909,Monrovia, [6°18’N, 10°48’W], 21 Oct. 1923, fl.(A!).NIGERIA: State Akwa Ibom: Talbot & Talbot 3364,Eket, [4°39’N, 7°56’E], 1912-1913, fl. (BM!). —State Cross River: Van Meer 1446, Oban Group ForestReserve, Kwa River, 150 m, [5°09’N, 8°28’E], 24 Apr.1971, fr. (WAG[2]!).REPUBLIC OF THE CONGO (Brazzaville): RegionBouenza: Sita 2460, région de Mouyondzi, environschutes de la Bouenza, [3°54’S, 13°46’E], 8 Jan. 1968,fr. (BR[2]!). — Region Kouilou: Dechamps 13155, LesSara, [4°22’S, 12°22’E], 28 June 1989, fl., fr. (BR!,MO!, WAG!). — Region Pool: Attims 172, Mayombe,15 km N du campement forestier de Ngongo, [4°41’S,14°32’E], 27 Mar. 1969, fr. (BR!).

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100 ADANSONIA, sér. 3 • 2005 • 27 (1)

2b. Octolepis casearia Oliv. var. flamignii (DeWild.) Z.S.Rogers, comb. et stat. nov.

Octolepis flamignii De Wild., Ann. Mus. CongoBelge, Bot., sér. 5, 3: 117, pl. 28. (1909). — Type:Flamigni 194A [cited as “Flamigni 194”], DemocraticRepublic of the Congo (Kinshasa), Prov. Kasaï-Oriental, Bene Dibele, [4°07’S, 22°50’E], 22 July 1907,fl., fr. (lectotype-, BR!, here designated; iso-, BR!).

Shrubs to 4 m tall, rarely trees to 7 m tall; youngstems densely strigose-tomentose. Leaf blades c. 5-22 × 3.5-7.6 cm, apex acuminate or acute, acumenup to 1.1 cm long. Inflorescence axes to 4 mmlong, with up to 14 flowers per inflorescence;pedicels 3-8 mm long. Flowers with 5 sepals. Fruitsdensely pubescent, surface obscured by dense per-sistent indument; trichomes to 0.5 mm long.

Refer to DE WILDEMAN (1909: 117, pl. 28) foran illustration of O. casearia var. flamignii.

DISTRIBUTION AND PHENOLOGY. — Octolepiscasearia var. flamignii occurs in the DemocraticRepublic of Congo (Kinshasa), the Republic ofthe Congo (Brazzaville), and southeastern Gabon(Fig. 4). The estimated elevational range for thespecies is 300-650 m. Octolepis casearia var.flamignii grows in primary, secondary and galleryforest, and sometimes on sand. It probably flow-ers and fruits year round. Flowers and fruits mayoccur simultaneously on a plant.

VERNACULAR NAMES. — Mukulo (Dechamps[R. 47] 100).

CONSERVATION STATUS. — Octolepis caseariavar. flamignii appears to be relatively widespreadin disturbed forests. Thus, the variety should beconsidered a taxon of Least Concern (LC).

TYPIFICATION. — Two sheets of the type col-lection (Flamigni 194A) are deposited at BR. A“TYPUS” label is affixed to both specimens, butonly one of them is fertile and annotated by DE

WILDEMAN. This particular sheet must have beenthe basis of the description and is designated asthe lectotype.

Octolepis casearia var. flamignii is easily distin-guished from var. casearia by the dense, persistentindument on the fruits (vs glabrescent or sparselystrigose fruits).

ADDITIONAL MATERIAL EXAMINED. — DEMO-CRATIC REPUBLIC OF THE CONGO (Kinshasa):Prov. Bandundu: Lebrun 6441, entre Kole et Bekese,Lac Leopold II, [2°00’S, 18°20’E], Oct. 1932, fl.(BM!, BR!); Vanderyst 3072, Kikwit, [5°02’S,18°49’E], Jan. 1914, fl. (BR!); Vanderyst 9723, entreIpamu et la Kamtsha, [3°42’S, 18°56’E], July 1921, fr.(BR!); Vanderyst 10085, same locality, July 1920, fr.(BR!); Vanderyst 10228, Mpio-Mpio, [3°52’S,16°05’E], Apr. 1921, fl. (BR!); Vanderyst 10240, samelocality, Aug. 1920, fr. (BR!). — Prov. Bas-Zaïre:Laurent s.n., Kole, Tsaka, [5°42’S, 16°04’E], 24 Nov.1903, fl. (BR!). — Prov. Équateur: Evrard 2812,Monkoto, Iwama, [2°08’S, 21°34’E], 12 Oct. 1957, fl.(BR!, MO!, WAG!); Evrard 5181, Ikela, Yongo,[0°17’S, 20°48’E], 19 Nov. 1958, fl. (BR!); Evrard5192, Ikela, Yalikunga-Yalomboka, [0°40’S, 22°41’E],20 Nov. 1958, fr. (BR!); Evrard 5493, Ikela,Mondombe, Bokone, [0°51’S, 22°36’E], 8 Jan. 1959,fr. (BR!); Germain 8356, Bokote, route Boende-Ingende, village Bumbanda, [0°44’S, 19°12’E],18 May 1954, fl., fr. (BM!, BR!). — Prov. Haut-Zaïre:Lisowski 43100, Haut-Zaïre, Zone d’Opala, près duvillage Lifera, entre Kaneke et Masua, [0°58’S,23°58’E], 14 Nov. 1976, fl. (BR!); Lisowski 43131,same locality, fl. (BR!). — Prov. Kasaï-Occidental:Dechamps (R. 47) 100, Kasai, Mweka, Kakenge,585 m, [04°51’S, 21°34’E], 16 Jan. 1959, fl. (BR!, K!).— Prov. Kasaï-Oriental: Claussens 208, Bene Dibele,[4°07’S, 22°50’E], Oct. 1904, fl. (BR(2)!); Luja 135,Sankuru, [4°58’S, 23°27’E], Dec. 1903, fl., fr. (BR!);Vankerckhoven 17, Sainte Trudon (= Mombelaye),[5°04’S, 23°29’E], June 1911, fl., fr. (BR!). —Kinshasa City: Breyne 799, Zone: Kinshasa, NdjiliAerodrome, [4°23’S, 15°27’E], 5 Mar. 1970, fl., fr.(BR!); Breyne 2345, same locality, 7 Mar. 1975, fl.(BR!); Muambi 11, Maluku, Plaine de Kinshasa,[4°18’S, 15°18’E], 9 Dec. 1967, fr. (BR!); Muambi 27,same locality, 5 Jan. 1968, fr. (BR!); Pauwels 4944,Léopoldville, Nkamu, Bingi-Bingi, [4°18’S, 15°18’E],30 Mar. 1965, fl. (BR!, MO!, WAG!).GABON: Prov. Haut-Ogooué: Breteler 6437, Moanda-Franceville, 1°39’S, 13°17’E, 12 Sept. 1970, fr. (MO!,WAG[2]!); Breteler 6738, Moanda-Bakoumba, km 30,1°45’S, 13°05’E, 7 Mar. 1975, fl. (BR!).REPUBLIC OF THE CONGO (Brazzaville): RegionBouenza: Sita 2669, Plateau des Cataractes, pisteTaba-Mandzakala, région de Kinkala, [4°27’S,13°52’E], 11 Oct. 1968, fl., fr. (BR[2]!).

3. Octolepis dioica Capuron

Adansonia, n.s. 3: 138 (1963). — Type: ServiceForestier (Capuron) 18002, Madagascar, Prov.Antananarivo, forêt Ambohitantely, sur le Tampoketsad’Ankazobe, c. 1600 m, [18°09’S, 47°18’E], June1957, fl. (holo-, P!; iso-, G!, K!, MO!, TEF!).


101ADANSONIA, sér. 3 • 2005 • 27 (1)

Octolepis dioica Capuron f. macrocarpa Capuron,Adansonia, n.s. 3: 140 (1963). — Type: ServiceForestier (Ravelojaona) 6028, Madagascar, Prov.Antananarivo, région de Tsinjoarivo (Amba-tolampy), [1500 m], [19°38’10”S, 47°41’30”E],27 Oct. 1952, fr. (holo-, P!; iso-, TEF!); syn. nov.

Trees to 20 m tall; young stems strigose-tomentose. Leaves alternate or subopposite;blades broadly obovate or elliptic, rarely subor-bicular, 2.0-6.0(-6.5) × 1.4-4.0 cm, length/width ratio c. 1.5-2.5:1, coriaceous, surfacesusually discolorous, adaxial surface glabrous,abaxial surface lighter green-brown, sparselystrigose or glabrescent, apex slightly emarginateto retuse, rarely acute, rounded or rarely shortcuspidate, margin strigose or glabrescent, revo-lute, more so near base, base short attenuate orcuneate; midrib strigose or glabrescent; venationstrongly raised to nearly inconspicuous on bothsurfaces, secondary veins c. 5-7 pairs, straight orslightly arched towards apex, angle of divergencefrom midrib 45-65°, submarginal vein loopc. 1-3 mm from margin; petioles 4-7 mm long,c. 1 mm in diam., strigose-tomentose or glabres-cent. Inflorescences axillary, usually borne ondefoliated portion of stem; fascicles 3-7-flowered,1 or 2 flowers persistent; pedicels 4-10(-14) mmlong, c. 0.5 mm in diam., glabrescent or sparselystrigose. Flower buds 2-3.5 × 2.5-3.5 mm,glabrescent, rarely sparsely strigose, ± smooth.Flowers 8-11 mm wide; sepals 4-5(6), ovateor oblong, rarely suborbicular or subtriangular,4-7(-10) × 2-4 mm, coriaceous, glabrescent onboth surfaces, apex acute or rounded, tomentose,margins glabrescent or tomentose; petals 4-5(6),densely to moderately strigose, rarely glabres-cent near base; each petal lobe narrowly trian-gular, (3.5-)4.5-6.5(-8.0) × 0.5-1.2 mm,length/width ratio c. 5-7:1, coriaceous, apexacute. Staminate flowers with spreading sepals;stamens 10; filaments 2.5-4.7 × 0.2-0.3 mm;anthers 1.5-2.4 × 0.8-1.3 mm; pisti l lodeminute, obscured by dense erect receptacle tri-chomes, trichomes 0.5-1.0 mm long. Pistillateflowers with erect sepals; staminodes 10, 0.5-1.2 mm long, 0.1-0.3 mm wide (at base),smooth or rugulose, glabrous; rudimentaryanthers globose or undifferentiated; ovary 1.8-3.4 × 1.8-2.4 mm, surrounded by many erect

receptacle trichomes, trichomes 0.7-0.9 mmlong; style 0.0-1.1 × 0.4-0.5 mm, glabrescent.Fruits ovoid-pyramidal, 1.9-3.0 × 1.6-2.3 cm;dehiscence lines 4 or 5 (i.e. 4 or 5 carpels deve-lop), usually strongly depressed, bicarinate onboth sides of line; pericarp fleshy, c. 4-6 mmthick, rugose or rugulose, sparsely or moderatelycovered with short indument, trichomes c. 1.0-1.5 mm long, subadpressed or erect; persistentstyle to 2 mm long. Seeds dark brown or black,1.3-1.9 cm long, 5-7 mm wide, puberulent.

Refer to CAPURON (1963: 139, pl. 2, Nos 1-10) for an illustration of O. dioica.

DISTRIBUTION AND PHENOLOGY. — Octolepisdioica occurs in humid forests along the centralplateau of Madagascar from c. 850-1650 melevation (Fig. 8). It has been collected as farnorth as Marojejy National Park, and as far southas Andohahela National Park (Parcel 1). Thespecies flowers from February to December andfruits from September to February. Floweringtime may be no more than one or two weeks inlength based on my observations of the popula-tion growing at Ranomafana National Park.Flowers and fruits do not occur on the same plantsimultaneously.

VERNACULAR NAMES . — Avoha (ServiceForestier 17898); Gavoala (Service Forestier34058); Fanamoratrakoho (Service Forestier17898); Fanamosatrakora (Service Forestier34480); Hafotramaladia (Service Forestier s.n.[20 Dec. 1951]); Havoa (Service Forestier 48-R-121, Service Forestier 6028); Valolahy (ServiceForestier 16804, 16824, 18001, 18002).

CONSERVATION STATUS. — Octolepis dioica isrelatively widespread in Madagascar and occursin several protected areas: Ambohitantely SpecialReserve, Andohahela National Park, MarojejyNational Park, Ranomafana National Park,Zahamena National Park. The extent of occur-rence of the species is c. 90000 km2 and the areaof occupancy is c. 80000 km2 based on a 100 kmgrid cell size (cell width/height, not area). At thepresent time, O. dioica should be considered aspecies of Least Concern (LC), but its status mayneed to be upgraded in the future, as suitablehabitat continues to disappear in the high plateauregion of Madagascar.

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102 ADANSONIA, sér. 3 • 2005 • 27 (1)

TYPIFICATION. — Two collections of Octolepisdioica were made from Ambohitantely SpecialReserve by René CAPURON in June 1957, one wasstaminate (Service Forestier 18001), while the otherwas pistillate (Service Forestier 18002). Specimensof both collections are deposited at P and are anno-tated with the word “Type” in CAPURON’s hand-writing, but only the latter was designated as thetype in the original publication, so it is thus to beregarded as the holotype. SF 18001 is a paratype.

Octolepis dioica is distinguished from O. ibi-tyensis, the most morphologically similar species,by a number of vegetative, floral, and fruit char-acters (Table 2). Octolepis dioica differs fromO. oblanceolata, a species previously described asa form of O. dioica, by its shorter leaves (longestleaf blades 2.8-6.0(-6.5) vs (7.0-)7.4-12.4 cmlong) with flat (vs undulate) margins and by itssmaller flowers (8-11 vs 11-13 mm wide).

CAPURON (1963) based Octolepis dioica f. macro-carpa on a single collection (Service Forestier 6028),which had smaller leaves and larger fruits withmore pronounced carinate dehiscence lines thanthe other specimens available to him at the time ofdescription. I observed overlapping variation inthese characters at Analamazoatra Special Reserveand Ranomafana National Park, which suggeststhat the variation in fruit morphology noted byCAPURON is most likely due to differences in matu-rity and the number of developed seeds.

Three collections are excluded from the descrip-tion of Octolepis dioica (Andrianjafy 209, ServiceForestier 28811, 28813). Both Service Forestier col-

lections, determined as O. dioica by CAPURON, weremade in 1969 from Kalalao forest, a small degradedforest on laterite on Île Ste Marie. Andrianjafy 209was recently collected at 600 m elevation inZahamena National Park. All three collections haveleaves similar to those found in O. dioica, but differby their densely sericeous pedicels and sepals.

ADDITIONAL MATERIAL EXAMINED. — MADAGAS-CAR: Service Forestier s.n. without precise locality, st.(TEF!). — Prov. Antananarivo: Rogers et al. 118,Ambohitantely Special Reserve, 2 km NE of the parkentrance, dense humid forest on laterite, 1600 m,18°10’08”S, 47°16’40”E, 6 Feb. 2003, st. (MO);Rogers et al. 119, same locality, st. (MO!); Rogers et al.123, same locality, st. (MO!); Rogers et al. 125, samelocality, st. (MO!); Service Forestier (Ravelojaona) 48-R-121, Tsinjoarivo, (Ambatolampy), [1500 m],[19°37’30”S, 47°41’30”E], 20 July 1951, st. (TEF!);Service Forestier (Ecol. Forest.) 16804, Manankazo-Ankazobe SF, village le plus proche Poste ForestierManankazo, [1300-1600 m], [18°09’00”S,47°14’00”E], 22 June 1955, y.fl. (TEF!); ServiceForestier (Ecol. Forest.) 16824, same locality, 22 June1955, y.fl. (P!, TEF[2]!); Service Forestier (Capuron)18001, Forêt d’Ambohitantely, sur le tampoketsad’Ankazobe, c. 1600 m, [18°09’30”S, 47°18’E], June1957, fl. (P[2]!, TEF[2]!). — Prov. Antsiranana:Rakotomalaza et al. 853, Marojejy Réserve NaturelleIntégrale, 10.5 km NW of Manantenina, along tribu-tary at head of Andranomifototra River, Camp 4,1625 m, 14°26’24”S, 49°44’30”E, 9 Nov. 1996, fr.(G!, MO!, P!, TAN!); Rakotonasolo & Ravelonarivo600, same locality, 1580-1650 m, 25 Feb. 2003,

y.fl. (MO!); Rakotonasolo & Ravelonarivo 601, samelocality, y.fl. (MO!). Prov. Fianarantsoa: Ravo-lolonanahary et al. 7, Ranomafana National Park, pistetouristique de Vohiparara, à l’Est du village deVohiparara, 1020-1170 m, 21°14’S, 47°23’E, 8 Oct.


103ADANSONIA, sér. 3 • 2005 • 27 (1)

TABLE 2. — Morphological differences between Octolepis dioica Capuron and O. ibityensis Z.S.Rogers.

Character Octolepis dioica Octolepis ibityensis

Leaf blade shape broadly elliptic or obovate (suborbicular) narrowly elliptic or obovateWidth of widest leaf blade (mm) (1.6-)2.0-4.0 1.0-1.4No. of flowers per inflorescence 3-7 1Pedicel length (mm) ≥ 4 ≤ 2Flower width (mm) 8-11 4-6Sepal length (mm) ≥ 4 2.5-3.5Petal lobe dimensions (mm) ≥ 3.5 × 0.5-1.2 2.1-3.2 × 0.4-0.5Filament length (mm) ≥ 2.5 1.6-2.0Anther dimensions (mm) 1.5-2.4 × 0.8-1.3 0.5-0.8 × 0.4-0.6Length of longest fruits (cm) 2-3 1.4-1.6No. of dehiscence lines on fruit 4 or 5 1 or 2(-3)Pericarp thickness (mm) ≥ 4 2-3

1996, fr. (MO!, P!, TAN!); Rogers et al. 63,Ranomafana National Park (Parcel 1), along piste A,1140 m, 21°14’12”S, 47°23’47”E, 17 Jan. 2003, fr.(MO!, P!, TAN!); Rogers et al. 64, same locality, fr.(MO!, P!, TAN!); Rogers et al. 65, same locality, fr.(MO!, P!, TAN!); Rogers et al. 66, same locality, st.(MO!). — Prov. Toamasina: Andrianjafy et al. 144,Parc National de Zahamena, près compound d’An-dalentitra, 1240-1400 m, 17°52’47”S, 48°44’21”E,28 Sept. 2001, fr. (MO!); Dorr & Barnett 3190,Analamazoatra Réserve Spéciale, [18°56’S, 48°25’E],2-5 Nov. 1984, fr. (MO!, TAN!); Perrier de la Bâthie4675, Forêt d’Analamazoatra, [18°56’S, 48°26’E], fr.(P[2]!); Razafitsalama et al. 491, Parc NationalMantadia, [18°50’S, 48°28’E], 27 Oct. 2003, fr.(TAN!); Razakamalala 748, Analamazoatra RéserveSpéciale, 1000 m, 18°56’12”S, 48°25’10”E, Sept.2003, fl. (MO!, P!, TAN!); Rogers et al. 46, same local-ity, 1000 m, 18°56’12”S, 48°25’10”E, 10 Jan. 2003,fr. (MO!, P!, TAN!); Rogers et al. 169, same locality,26 Feb. 2003, st. (MO!); Rogers et al. 170, same local-ity, 26 Feb. 2003, st. (MO!); Rogers et al. 171, samelocality, 26 Feb. 2003, st. (MO!); Service Forestier s.n.,Analamazoatra-Perinet, S.P. 36, 20 Dec. 1951, st.(TEF!); Service Forestier (Raderasolo) 4411, same local-ity, 15 Oct. 1951, fl. (TEF!); Service Forestier (Gachet)17898, same locality, 20 Feb. 1958, fr. (P!, TEF!);Service Forestier (Capuron) 18052, same locality, 29-30 July 1957, y.fl. (P!, TEF!); Service Forestier(Capuron) 24346, same locality, 21 Dec. 1965, fr. (K!,MO!, P!, TEF!); Service Forestier 34058, FiraisanaAndasibe, Fivondronona Moramanga, Ampan-galatsary, crête Nord, [1000 m], [18°56’S, 48°26’E],6 Sept. 1990, fl., fr. (MO!, TEF!); Service Forestier(Comtet) 34480, Fokontany Faliarana, FiraisanaAndasibe, Fivondronona Moramanga, village le plusproche Faliarana, au bord de la rivière de Iofa,[18°47’S, 48°33’E], 15 Oct. 1993, fr. (TEF[2]!). — Prov.Toliara: Messmer et al. 114, Andohahela NationalPark, parcelle 1, 15 km NW of Eminiminy, 1500 m,24°34’10”S, 46°43’55”E, 24 Nov. 1995, fr. (G!, MO!).

4. Octolepis ibityensis Z.S.Rogers, sp. nov.

Haec species a Octolepide dioica, cui aliter similis est,foliis angustioribus 0.8-1.4 (haud 1.4-4.0) cm latis,inflorescentiis unifloris (haud 3- ad 7-floris), pedicellisminus quam 2 (haud plus quam 4) mm longis et fructu1.4-1.6 cm longo lineis dehiscentiae 1 vel 2 (haud 1.9-3 cm longo lineis (4 ad 5)) differt.

TYPUS. — Randrianaivo, Ranaivojaona, Birkinshaw,Ravololonanahary, Ralimanana & Randrianasolo 8,Madagascar, Prov. Antananarivo, Mont Ibity, à 2-3 km à l’Est de la Cimenterie d’Ibity, 1590-1830 m,20°04’40”S, 47°00’10”E, 19-21 Oct. 1993, fl.(holo-, MO!; iso-, G!, P, TAN!, WAG!).

Trees to 3 m tall; young stems sparsely moder-ately to sparsely strigose-tomentose or glabres-cent. Leaves alternate to subopposite, rarelyopposite; blades narrowly obovate or elliptic,c. 2.6-4.6 × 0.8-1.4 cm, length/width ratio c. 3-5:1, coriaceous, surfaces discolorous, adaxial sur-face glabrescent, abaxial surface lighter green,sparsely strigose or glabrescent, apex emarginate,usually with a small notch, rarely rounded, mar-gin sparsely strigose or glabrescent, revolute,more so near base, base short attenuate orcuneate; midrib sparsely strigose or glabrescent;venation strongly raised on both surfaces, sec-ondary veins c. 6-7 pairs, ± parallel, straight orarched toward apex, angle of divergence frommidrib 30-55°, submarginal vein loop c. 1-2 mmfrom margin; petioles 3-5 mm long, c. 1 mm indiam., strigose-tomentose or glabrescent.Inflorescences axillary, borne on the defoliatedportion of the stem; fascicles 1-flowered; pedicelsc. 1-2 mm long, c. 1 mm in diam., glabrescent orsparsely strigose. Flower buds 2.0-2.5 × 2.0-2.5 mm, sparsely strigose, ± smooth. Flowers 4-6 mm wide; sepals 5, rarely 6, ovate, c. 2.5-3.5 ×2.0-2.5 mm, coriaceous, strigose-tomentose orglabrescent on both surfaces, ± smooth, apexacute or rounded, margin tomentose; petals 5,rarely 6, moderately to sparsely pubescent onboth surfaces; each petal lobe narrowly triangular,2.1-3.2 × 0.4-0.5 mm, length/width ratio c. 5-8:1, coriaceous, apex acute. Staminate flowerswith 10 stamens, rarely 12; filaments 1.6-2.0 ×0.2-0.3 mm; anthers 0.5-0.8 × 0.4-0.6 mm; pis-tillode minute, obscured by dense erect receptacletrichomes, trichomes c. 0.5-0.8 mm long.Pistillate flowers (description based on persistentorgans in fruit) with 10 staminodes, rarely 12;staminodes 0.8-1.2 mm long, 0.2-0.4 mm wide(at base), ± smooth, glabrous; rudimentaryanthers globose; ovary surrounded by many erectreceptacle trichomes, trichomes c. 0.5-0.8 mm long.Fruits ovoid or ovoid-pyramidal (with 3 faces),1.4-1.6 × 1.2-1.4 cm; dehiscence lines 1 or 2,rarely 3 (i.e. 1-3 carpels develop), depressed, notbicarinate; pericarp fleshy, c. 2-3 mm thick,rugose or rugulose, sparsely strigose to nearlyglabrous, trichomes of different lengths, tri-chomes 0.7-1.0 mm long or 0.1-0.2 mm long;persistent style to 1 mm long. Seeds dark brown

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104 ADANSONIA, sér. 3 • 2005 • 27 (1)


105ADANSONIA, sér. 3 • 2005 • 27 (1)

A B

C

D

0.5 cm

0.5 cm

1.5 cm

1 cm

FIG. 5. — Octolepis ibityensis Z.S.Rogers, sp. nov.: A, habit; B, staminate flower; C, relationship between sepals, bifid petals, andstamens; D, fruit. A, D, Rogers et al. 80 (TAN); B, C, Randrianaivo et al. 8 (TAN, type).

or black, 1.1-1.2 cm long, 5-6 mm wide, sparselypuberulent, more evident near the poles. — Fig. 5.

DISTRIBUTION AND PHENOLOGY. — Octolepisibityensis grows in narrow tracts of gallery forestalong two ravines on the Ibity Massif in centralMadagascar from 1450 to 1830 m elevation(Fig. 8). The species flowers from October toJanuary and fruits in January.

VERNACULAR NAME. — None available.CONSERVATION STATUS. — Octolepis ibityensis

is known from two remnant gallery forests on theslopes of the unprotected Ibity Massif. The lowerslopes of the mountain are burned annually bylocal villagers, leaving only narrow (c. 5-15 mwide) tracts of forests running along streams thatfunction as natural firebreaks and provide enoughshelter to protect the small populations of O. ibi-tyensis growing along them. However, due to thefrequent burning, the tracts continue to narrow,and plants untouched by fire in 2003, have sincebeen damaged to the point that they may notrecover. Octolepis ibityensis is given the provisionalstatus of Critically Endangered (CE B1ab+B2ab).

Octolepis ibityensis is distinguished from O. dioica,the most morphologically similar species, by itsnarrowly elliptic leaf blades, its shorter pedicels, itsfewer flowered inflorescences, its smaller flowers(with shorter sepals, smaller petal lobes, shorter fila-ments, and smaller anthers), and its smaller fruitswith fewer lines of dehiscence. Table 2 summarizesmorphological differences between both species.

PARATYPES. — MADAGASCAR: Prov. Antana-narivo: Rogers et al. 78, Ibity massif, c. 2-3 km E of theIbity cement factory, along mountain stream of Be-Apombo, 1540 m, 20°04’14”S, 47°00’11”E, 20 Jan.2003, st. (MO!); Rogers et al. 79, same general local-ity, along mountain stream of Falirano, 1470 m,20°04’24”S, 47°00’10”E, 20 Jan. 2003, y.fl.(MO!); Rogers et al. 80, same locality, fr. (MO!, P!,TAN!); Rogers et al. 81, same locality, st. (MO!); Rogerset al. 306, same locality, 28 Apr. 2004, st. (MO!).

5. Octolepis oblanceolata (Capuron) Z.S.Rogers,comb. et stat. nov.

Octolepis dioica Capuron f. oblanceolata Capuron,Adansonia, n.s. 3: 140 (1963). — Type: Réserves

Naturelles (Rakotoson) 10459, Madagascar, Prov.Toliara, Marovato, District: Amboasary, [24°41’S,46°45’E], 16 June 1960, fl. (lecto-, P-00077575!, here designated; iso-, P[2]!, TEF!).

Trees to 10 m tall; young stems moderately tosparsely strigose-tomentose or glabrescent. Leavesalternate to opposite; blades broadly to narrowlyobovate, rarely broadly elliptic, c. 6.5-12.4 × 1.3-4.0 cm, length/width ratio c. 2-6:1, coriaceous,surfaces discolorous, adaxial surface glabrous,abaxial surface lighter green-brown, sparselystrigose or glabrescent, apex usually acute, some-times emarginate or short cuspidate, marginsparsely strigose or glabrescent, revolute, undu-late, base attenuate or cuneate; midrib strigose orglabrescent; venation raised to nearly inconspi-cuous on both surfaces, secondary veins c. 8-12 pairs, ± parallel, arched toward apex in upperhalf of leaf, angle of divergence from midrib 35-55°, submarginal vein loop c. 2-3 mm from mar-gin; petioles 4-8(-9) mm long, c. 1.0-1.5 mm indiam., glabrescent or sparsely strigose. Inflo-rescences axillary, usually borne on foliated por-tion of the stem; fascicles 2-flowered, usually1 flower persistent; pedicels (0.9-)1.1-1.5 cmlong, 0.5-0.7 mm in diam., glabrescent or rarelysparsely strigose. Flower buds 3.5-4 × 3.5-4 mm,glabrescent, ± smooth. Flowers 1.1-1.3 cm wide;sepals 4 or 5, ovate or oblong, rarely subtriangu-lar, 5-6(-8) × 2-3(-4) mm, coriaceous, glabres-cent, rarely sparsely strigose on both surfaces,apex acute or rounded, tomentose, margins gla-brescent or tomentose; petals 5, rarely 6, denselyto sparsely strigose on both surfaces; each petallobe narrowly triangular, 3.8-5.2 × 0.8-1.1 mm,length/width ratio 4-5:1, coriaceous, apex acute.Staminate flowers with 10 stamens, rarely 12; fil-aments c. 3.5-4.5 × 0.3-0.4 mm; anthers c. 2 ×1 mm long; pistillode minute, obscured by denseerect receptacle trichomes, trichomes c. 1 mmlong. Pistillate flowers (description based on per-sistent organs in fruit) with 10 (12) staminodes;staminodes 1.3-1.9 mm long, 0.3-0.4 mm wide(at base), smooth; rudimentary anther globose;ovary surrounded by many erect receptacle tri-chomes, trichomes c. 2 mm long. Fruits sub-spheroidal or ovoid-pyramidal, 1.7-2.1 ×1.7-2.4 cm; dehiscence lines 4 or 5 (i.e. 4 or5 carpels develop), ± bicarinate on both sides of

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106 ADANSONIA, sér. 3 • 2005 • 27 (1)

line; pericarp fleshy, 4-5 mm thick, rugulose ornearly smooth, sparse or moderately pubescent,trichomes c. 1-2 mm long, erect or subadpressed,± rigid, not stinging; persistent style to 4 mmlong. Seeds dark brown, 1.1-1.3 cm long, 5-6 mm wide, puberulent, indument denser nearpoles.

Refer to CAPURON (1963: 139, pl. 2, No. 12)for an illustration of O. oblanceolata.

DISTRIBUTION AND PHENOLOGY. — Octolepisoblanceolata occurs in southeastern Madagascarnear Fort-Dauphin from c. 400-700 m elevation(Fig. 7). The species has been collected inAndohahela National Park and was also found in2003 growing on the forested slopes aboveSt Jacques, a small village overlooking Fort-Dauphin. Since then, however, the population atSt Jacques has been extirpated (J. RABENANTO-ANDRO pers. comm.). The species flowers in Juneto November and fruits from August to January.

VERNACULAR NAME. — Tsilorano (RéservesNaturelles 10072).

CONSERVATION STATUS. — Octolepis oblanceo-lata is known from three or four populations,perhaps only one of which is formally protectedwithin the boundaries of Andohahela NationalPark (Parcel 1). The population above St Jacquesdiscovered in 2002 has recently been lost. Localitydata for the other collections are incomplete,making it difficult to calculate extent of occur-rence and area of occupancy. The species isassigned a preliminary conservation status ofEndangered (EN B1ab+B2ab).

TYPIFICATION. — Three sheets of the type col-lection (Réserves Naturelles 10459) are depositedat P. Each of the specimens was identified astype of Octolepis dioica f. oblanceolata byCAPURON. The sheet, bearing the accession num-ber P-00077575, was annotated as the “Type dela forme”, the second (P-00077576) bears an iso-type label affixed to the lower half of CAPURON’sannotation slip, and the third is marked “Isotyped. l. [= de la] forme”. The style of the labels, thehandwriting, and the completeness of the labelinformation differs between all three sheets, butnone of the data on the labels contradicts the pro-tologue. Sheet (P-00077575) is the only sheet

without the isotype designation and is selected asthe lectotype.

Octolepis oblanceolata was originally consideredto be a form of O. dioica by CAPURON (1963). Thespecies is distinguished from O. dioica by its larger,oblanceolate or more broadly obovate leaves, itsmore strongly arched secondary veins in the upperhalf of the leaves, its more often acute apices, itsundulate leaf margins, its longer pedicels ((9.0-)11.0-15.0 vs 4.0-10.0(-14.0) mm long), and itslarger flowers (11-13 vs 8-11 mm wide).

Four collections are excluded from Octolepisoblanceolata as circumscribed above. Three ofthem (A. Randrianasolo 790, Ludovic 721 and734) have been made recently at Mahabo forest, alittoral forest fragment along Madagascar’s south-east coast, and appear to be the first ever collectedon sand in Madagascar (Octolepis casearia is fre-quently collected on sand in continental Africa).The fourth collection, Service Forestier 25524,was made in 1965 with an incomplete locality onthe label, although it was probably made some-where to the south of Mantadia National Park.There is also a good possibility that the collectionwas made near the coast, because it shares mor-phological similarities with the Mahabo speci-mens. All four collections have leaves that areclearly larger than those of O. dioica, resemblingsome of the broadly oblanceolate leaves found onplants of O. oblanceolata. However, theseexcluded specimens differ in having denselysericeous pedicels, and their smaller, much moredensely pubescent fruits with 3, or rarely 4,dehiscence lines.

ADDITIONAL MATERIAL EXAMINED. — MADAGAS-CAR: Prov. Toliara: Randriamampionona 1308,Andohahela National Park, Emalio, Andranolava, Sitede suivi éco-tourisme, [24°41’S, 46°45’E], 25 Aug.1996, fr. (MO!, P, TAN!); Réserves Naturelles(Rakotoson) 10072, Canton Behara, districtAmboasary, [24°41’S, 46°45’E], 5 Nov. 1958, fl.(P!, TEF!); Randrianaivo et al. 880, FivondrononaFort-Dauphin, Commune (Fokontany) Ampasinampoa,Forêt de Ambatomitikitra, à 3 km W du villageSt Jacques, forêt humide de basse altitude, 400-700 m,24°58’56”S, 46°57’30”E, 2 Dec. 2002, fr. (MO!, P,TAN!); Rogers et al. 101, same locality, 520-660 m,24°58’57”S, 46°57’24”E, 25 Jan. 2003, st. (MO!);Rogers et al. 102, same locality, fr. (MO!, P!, TAN!);Rogers et al. 103, same locality, fr. (MO!).


107ADANSONIA, sér. 3 • 2005 • 27 (1)

6. Octolepis ratovosonii Z.S.Rogers, sp. nov.

Haec species quoad speciebus aliis Octolepidis abpedicellis 1.6-2.5 cm longis, petalis 10 (haud 5), sta-minibus vel staminodiis 20 (haud 10, raro 8 vel 12) etpericarpio indumento denso bistrato praedito strato exte-riore trichomatibus urticantibus constante differt.

TYPUS. — Rogers, Rakotonasolo & Ravelonarivo 165,Madagascar, Prov. Antsiranana, Anjanaharibe-SudRéserve Spéciale, c. 2 km NE of main road, along trailto Ranomafana thermal spring, 976 m, 14°45’05”S,49°29’45”E, fr. (holo-, MO!; iso-, BR!, K!, P!, TAN!,WAG!).

Trees to 9 m tall; young stems densely tosparsely strigose-tomentose. Leaves alternate toopposite; blades broadly elliptic or obovate, c. 6.6-16.5 × 2.3-5.3 cm, length/width ratio c. 2.5-4:1,coriaceous, surfaces discolorous, dark green andglabrous adaxially, abaxial surface dries a distinc-tive light green, sparsely strigose or glabrescent,apex retuse or notched, margin sparsely strigoseor glabrescent, flat or slightly undulate, stronglyrevolute, more so near base, base short attenuateor cuneate; midrib moderately strigose or nearlyglabrous; venation raised on both surfaces, higherorder venation raised or inconspicuous, sec-ondary veins c. 7-11 pairs, angle of divergencefrom midrib 45-65°, submarginal vein loop c. 3-6 mm from margin; petioles 8-16 mm long, 1-2 mmin diam., strigose-tomentose. Inflorescences axil-lary, borne on foliated or defoliated portion ofstem, 1-flowered; pedicels (in fruit) 1.6-2.5 cmlong, (1.5-)2.0-3.0 mm in diam., surfaceobscured by dense indument. Flower buds 5.1-7.0 × 4.0-6.1 mm, surface obscured by denseindument, rugose. Flowers (description based onstaminate flower buds and persistent organs infruit) with 4-6 sepals; sepals triangular or ovate,(0.8-)1.0-1.1 × 0.5-0.9 cm, very coriaceous, c. 2-4 mm thick (in fruit), both surfaces obscured bydense indument, trichomes c. 0.3-0.4 mm long;petals 10, densely to moderately strigose; eachpetal lobe narrowly triangular, 2.4-3.1 × 0.4-0.6 mm, length/width ratio c. 4-6:1, coriaceous,apex acute. Staminate flowers with 20 stamens.Pistillate flowers with 20 staminodes; staminodes1.5-1.9 mm long, 0.3-0.4 mm wide (at base),± smooth, glabrous; rudimentary anthers well

developed, c. 1.0-1.1 × 0.3-0.4 mm; ovarysurrounded by many erect receptacle trichomes,t r i c h o m e s c . 1 . 5 - 2 . 5 m m l o n g . Fr u i t sovoid-subspheroidal, 2.4-2.9 × 2.2-2.8 cm;dehiscence lines (4-)5 or 6 (i.e. 4-6 carpelsdevelop), strongly depressed, bicarinate on bothsides of line; pericarp fleshy, 7-8 mm thick,rugose and pitted, surface obscured by 2-layeredindument, shorter layer dense, trichomes 0.2-0.3 mm long, erect, soft, longer layer sparse, tri-chomes c. 1.5 mm long, erect, stiff, stinging(when fresh); persistent style 2-4 mm long. Seedsred-brown, 1.3-1.7 cm long, 7-9 mm wide, sur-face obscured by dense puberulent indument. —Fig. 6.

DISTRIBUTION AND PHENOLOGY. — Octolepisratovosonii occurs in humid forests, from 800-1250 m elevation (Fig. 7). The species has beencollected in flower bud in February and in fruitin February, July, and August.

VERNACULAR NAMES. — None available.CONSERVATION STATUS. — Octolepis ratovosonii

is known from three formally protected areas(Anjanaharibe-Sud RS, Manongarivo RS,Zahamena PN). The extent of occurrence of thespecies is c. 28000 km2. Area of occupancy isc. 10000 km2 based on a 50 km grid cell size (cellwidth/height, not area). Octolepis ratovosoniishould be considered Near Threatened (NT).

Octolepis ratovosonii differs markedly from allother species in the genus by having 10 (vs 4-6)petals, 20 (vs 10, rarely 8 or 12) stamens and sta-minodes, and larger flowers (probably 3-4 timeslarger than other species based on the flower buds).Other distinguishing features include its denselypubescent, long and robust (c. 1.6-2.5 cm × 1.5-3.0 mm) pedicels, its large, thickened, densely pu-bescent sepals, its thick, rugose, pericarp coveredin by a two-layered indument, the longer of whichis composed of stinging trichomes, and its denselypuberulent seeds. Octolepis ratovosonii can berecognized vegetatively from the other Malagasyspecies by having the largest, thickest, leaves withthe most revolute margins, and by the light greencolor of the blades after drying.

The epithet has been chosen in honor ofFidisoa RATOVOSON, a Malagasy botanist, who

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108 ADANSONIA, sér. 3 • 2005 • 27 (1)


109ADANSONIA, sér. 3 • 2005 • 27 (1)

A

B C

D

1 cm 1 cm 1 cm 1 cm

FIG. 6. — Octolepis ratovosonii Z.S.Rogers, sp. nov.: A, habit; B, fruit capsule; C, capsule after dehiscence; D, densely puberulentseed. Rogers et al. 165 (TAN, type).

collected the species on two occasions atZahamena National Park, and whose contribu-tions have greatly increased our knowledge of theflora in that region.

PARATYPES. — MADAGASCAR: Prov. Antsi-ranana: Razafimandimbison et al. 229, Anjananharibe-Sud Réserve Spéciale, au sud de campement àMandritsarahely, 985 m, 14°46’S, 49°30’E, 10 July1996, fr. (G!, MO!, TAN!); Rogers et al. 164,Anjanaharibe-Sud Réserve Spéciale, c. 2 km NE ofmain road, along trail to Ranomafana thermal spring,976 m, 14°45’05”S, 49°29’45”E, 18 Feb. 2003, y.fl., fr. (MO!); Rogers et al. 166, same locality, y.fl.(MO!); Wohlhauser et al. 350, Manongarivo RéserveSpéciale, Vallée de l’Ambahatra (cours supérieur),Bassin de l’Ambahatra (bras gauche, à l’ouestd’Andetryfotsy), 1200 m, 14°00’S, 48°26’E, 21 Nov.

2000, fr. (G!). — Prov. Toamasina: Ratovoson et al. 247,Zahamena National Park, Fivondronana Ambaton-drazaka, Firaisana Antanandava, Ankosy, Ambarikely(3 km à l’Est d’Ankosy), 996-1250 m, 17°28’48”S,48°49’33”E, 12 July 2000, y.fr. (MO!, P, TEF!);Ratovoson et al. 726, Zahamena National Park, 800-850 m, 17°29’S, 48°43’E, 3 Aug. 2003, fr. (MO!, P, TEF!);Service Forestier (Razakamalala & Norbert) 34487,Ambodivoahangy, Fir. Antanandava, Fiv: Ambato-drazaka, [17°56’S, 48°26’E], 15 Sept. 1997, fr. (TEF!).

Acknowledgements

I would like to thank the curators of the followingherbaria for making material available for this study:A, B, BM, BR, F, G, GH, K, MA, MO, NY, P,S, TAN, TEF, US, WAG. Special thanks toK.J. WURDACK for furnishing unpublished molecular

Rogers Z.S.

110 ADANSONIA, sér. 3 • 2005 • 27 (1)

45°E 50°E

15°S

20°S

25°S

200 km

45°E 50°E

15°S

20°S

25°S

200 km

F IG. 7. — Distribution map of Octolepis aymoninianaZ.S.Rogers (▲), O. oblanceolata (Capuron) Z.S.Rogers (■), andO. ratovosonii Z.S.Rogers (✚).

FIG. 8. — Distribution map of Octolepis dioica Capuron (●) andO. ibityensis Z.S.Rogers (★).

data. L.R. ANDRIAMIARISOA and S. HIRTH providedthe illustrations. P.F. STEVENS and R.E. GEREAU trans-lated the diagnoses into Latin. H. BRETELER ,P.P. LOWRY II, P.B. PHILLIPSON and P.F. STEVENSprovided many helpful review comments on the man-uscript. Several other people should be acknowledgedfor var ious contr ibutions to this research:S. ANDRIAMBOLOLONERA, R. LUDOVIC, G. MCPHERSON,J. RABENANTOANDRO , J. RAHARIMAMPIONONA ,F. RAKOTONASOLO, A. RANDRIANASOLO, F. RATO-VOSON, R. RAZAKAMALALA, G.E. SCHATZ, P. SWEE-NEY, F. TRONCHET, and R. VOGT. Field work waspartially supported by the 2003 John DENVERMemorial Scholarship granted by the InternationalCenter for Tropical Ecology at the University ofMissouri-St Louis and by several grants to theMissouri Botanical Garden administered byP.P. LOWRY II and P.M. RICHARDSON.

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Submitted on 26 November 2004;accepted on 5 April 2005.


111ADANSONIA, sér. 3 • 2005 • 27 (1)

Documents

A revision of Octolepis Oliv. (Thymelaeaceae,Octolepidoideae)€¦ · species of Octolepis(with three forms) from Madagascar, named Octolepis dioicaCapuron; African Octolepisare monoecious